Immediate and delayed life history effects caused by food deprivation early in life in a short‐lived lizard

Detailed studies of the mechanisms driving life history effects of food availability are of prime importance to understand the evolution of phenotypic plasticity and the capacity of organisms to produce better adapted phenotypes. Food availability may influence life history trajectories through three nonexclusive mechanisms: (i) immediate and long‐lasting effects on individual quality, and indirect delayed effects on (ii) intracohort and (iii) intercohort interactions. Using the common lizard (Zootoca vivipara), we tested whether a food deprivation during the two‐first months of life influence life history (growth, survival, reproduction) and performance traits (immunocompetence, locomotor performances) until adulthood. We investigated the underlying mechanisms and their possible interactions by manipulating jointly food availability in a birth cohort and in cohorts of older conspecifics. Food deprivation had direct immediate negative effects on growth but positive long‐lasting effects on immunocompetence. Food deprivation had also indirect delayed effects on growth, body size, early survival and reproduction mediated by an interaction between its direct effects on individual quality and its delayed effects on the intensity of intercohort social interactions combined with density dependence on body size. These results demonstrate that interactions between direct and socially mediated effects of past environments influence life history evolution in size‐structured and stage‐structured populations.     

Adjustment of Parental Investment in the Dung Beetle Onthophagus atripennis (Col., Scarabaeidae)

If parents can invest resources optimally per offspring, they should adjust the amount of investment in an offspring according to environmental heterogeneity. Many studies have demonstrated changes in egg size or the amount of resource supplied in response to environmental heterogeneity. However, it remains unclear whether parents simply know the resource type a priori or can assess resource quality and adjust the quantity of investment accordingly. We examined the parental capability to adjust the amount of investment per offspring by providing Onthophagus atripennis dung beetle parents with one of three dung types of different quality: monkey dung (high quality), cow dung (low quality), or a mixture of monkey and cow dung (medium quality). The beetle parents cooperatively produce dung brood masses each with one egg under the ground. The size of a brood mass, on which a larva can only feed until adult, represents a large part of the amount of investment. Parents produced a greater number of smaller brood masses given high‐quality resource, while they compensated for low quality of the resource by providing a larger amount of the resource, at the cost of offspring number. However, despite this compensation in the amount of food, offspring raised on low‐quality food was still smaller than offspring raised on high‐quality food. Thus, O. atripennis parents assessed resource quality partly and adjusted the amount of resource provided for their offspring.     

Among‐individual heterogeneity in maternal behaviour and physiology affects reproductive allocation and offspring life‐history traits in the garter snake Thamnophis elegans

Accumulating evidence suggests that within‐individual plasticity of behavioural and physiological traits is limited, resulting in stable among‐individual differences in these aspects of the phenotype. Furthermore, these traits often covary within individuals, resulting in a continuum of correlated phenotypic variation among individuals within populations and species. This heterogeneity, in turn, affects individual fitness and can have cross‐generational effects. Patterns of trait covariation, among‐individual differences, and subsequent fitness consequences have long been recognized in reptiles. Here, we provide a test of patterns of among‐individual heterogeneity in behaviour and physiology and subsequent effects on reproduction and offspring fitness in the garter snake Thamnophis elegans. We find that measures of activity levels vary among individuals and are consistent within individuals in reproductive female snakes, indicating stable behavioural phenotypes. Blood hormone and glucose concentrations are not as stable within individuals, indicating that these traits do not describe consistent physiological phenotypes. Nonetheless, the major axes of variation in maternal traits describe behavioural and physiological phenotypes that interact to predict offspring body condition and mass at birth. This differential allocation of energy to offspring, in turn, strongly influences subsequent offspring growth and survival. This pattern suggests the potential for strong selection on phenotypes defined by behaviour–physiology interactions.     

Provisioning habitat with custom‐designed nest‐boxes increases reproductive success in an endangered finch

Manipulating resource availability to assess the strength and effects of resource limitation on an animal population is relatively straightforward, and thus, common in the scientific literature. Resource quality, however, is rarely manipulated, as this requires a priori knowledge of a relationship between some measurable variation in the resource and individual preference and/or fitness. Recent research on nest‐site selection was used to inform the design of custom‐built nest‐boxes for an endangered hollow‐nester, the Gouldian finch (Erythrura gouldiae). By provisioning breeding populations with these ‘high quality’ artificial nest sites over 3 years, we experimentally investigated the strength and effects of nest‐site limitation in this species. Breeding pairs using nest‐boxes initiated nesting earlier, produced larger clutches and fledged more offspring per season than those in natural hollows. Total and mean reproductive output also increased in nest‐box provisioned sites over the duration of the study. All of these effects were predominantly driven by pairs initiating clutches earlier, potentially because of the reduced intra‐ and interspecific competition for nest sites at optimal breeding times. Our findings suggest that reproduction in wild Gouldian finch populations may be limited by the availability of high quality nest sites in the landscape, and that nest‐boxes could be used as a tool for enhancing reproduction in recovering populations. Furthermore, we conclude that resource quality is equally, if not more important than quantity when supplementing resource‐limited habitats.     

Parental age and lifespan influence offspring recruitment: a long-term study in a seabird

Recent studies of wild populations provide compelling evidence that survival and reproduction decrease with age because of senescence, a decline in functional capacities at old ages. However, in the wild, little is known about effects of parental senescence on offspring quality. We used data from a 21-year study to examine the role of parental age on offspring probability of recruitment in a long-lived bird, the blue-footed booby (Sula nebouxii). Offspring probability of recruiting into the breeding population varied over the life of parents and effects age were similar in mothers and fathers. Offspring recruitment was high when parents were roughly 6-12 years old and low before and after then. Effects of parental age on offspring recruitment varied with lifespan (parental age at last reproduction) and previous breeding experience. Offspring recruitment from young and old parents with long reproductive lifespans was greater than that of offspring from parents with short lifespans at young and old ages. For parents with little previous breeding experience recruitment of offspring decreased with their hatch date, but experienced parents were no similarly affected. We found evidence of terminal effects on offspring recruitment in young parents but not in older parents, suggesting that senescence is more likely a gradual process of deterioration than a process of terminal illness. Failure to recruit probably reflects mortality during the first years after independence but also during the fledgling transition to full independence. Our results show effects of parental age and quality on offspring viability in a long-lived wild vertebrate and support the idea that wild populations are composed of individuals of different quality, and that this individual heterogeneity can influence the dynamics of age-structured populations.     

Costs of rearing and sex‐ratio variation in southern grey shrike Lanius meridionalis broods

Several non‐mutually exclusive hypotheses predict adaptive variation in the offspring sex ratio. When conditions for breeding are adverse, parents are predicted to produce more offspring of the less costly sex to rear (‘the cost‐of‐reproduction hypothesis’). Moreover, they also should produce the more dispersing sex in order to diminish future competition (‘the local‐resource‐competition hypothesis’). Here, we analyse brood sex ratio according to rearing conditions in the southern shrike Lanius meridionalis, a species with moderately reversed sexual dimorphism. Our results suggest that females are more costly to rear than males in this species. Adult females proved heavier than males, and female nestling tended to be heavier than male nestlings. Moreover, the greater brood reduction, the more male‐biased was the brood, suggesting that brood reduction implied higher mortality in female nestlings. Consistent with these findings, the brood sex ratio was biased to the less costly sex (males) when breeding conditions were adverse (bad years or low‐quality male parents), supporting the cost‐of‐reproduction hypothesis. By contrast, these findings did not support the local‐resource‐competition hypothesis, which predicted female‐biased brood sex ratio under adverse conditions. As a whole, our results support the idea that birds adaptively modulate sex ratio in order to minimize reproduction costs.     

Is fatter fitter? Body storage and reproduction in ten populations of the freshwater amphipod Gammarus minus

Relationships between body storage (estimated as fat content and residuals of body mass regressed against body length) and offspring investment [brood mass, brood size (number of embryos per brood) and embryo mass] were examined within and among populations of the amphipod Gammarus minus in ten cold springs in central Pennsylvania, USA. Two major hypotheses and six corollary hypotheses were tested. Total reproductive investment (brood mass and brood size) was usually strongly positively correlated with maternal body length and body storage both within and among populations. These positive associations between reproductive and somatic investments are expected if individual variation in resource acquisition exceeds that of resource allocation. That is, individuals or populations that are able to acquire more resources should also be able to allocate more resources to both reproduction and somatic reserves than those acquiring fewer resources. This hypothesis is consistent with evidence showing that individual differences in body storage in G. minus and other amphipods are related to differences in resource acquisition. Positive associations between reproductive and somatic investments do not mean that energy costs of reproduction do not exist in G. minus. Evidence for reproductive energy costs included the lower body-fat contents of brooding versus nonbrooding females and the relatively low body mass per length of females who had just deposited eggs in their brood pouch. Unlike brood mass and brood size, individual embryo mass was usually unrelated to maternal body length and body storage. This pattern is largely consistent with optimal offspring investment theory, which predicts that offspring size should be insensitive to variation in parental resource status. However, in contrast to theory, embryo mass increased in winter when brooding females were significantly ”fatter”, presumably due to the availability of autumn-shed leaf food. This seasonal change in offspring size may be a maternally mediated effect of increased resource availability, though other explanations are possible. Overall, this study suggests that ”fatter” female amphipods are fitter than ”thinner” ones, though both the costs and benefits of increased body storage and brood size require investigation to substantiate this claim. This study also suggests that effects of individual variation in resource acquisition on life-history patterns deserve more theoretical and empirical attention by ecologists than they have received. It should be recognized that positive and/or nonsignificant correlations between life-history traits are just as interesting and important as are the negative correlations predicted by many theoretical models. Received: 20 January 1999 / Accepted: 26 September 1999     

Telomere attrition and growth: a life‐history framework and case study in common terns

The relationship between growth and age‐specific telomere length, as a proxy of somatic state, is increasingly investigated, but observed patterns vary and a predictive framework is lacking. We outline expectations based on the assumption that telomere maintenance is costly and argue that individual heterogeneity in resource acquisition is predicted to lead to positive covariance between growth and telomere length. However, canalization of resource allocation to the trait with a larger effect on fitness, rendering that trait relatively invariant, can cause the absence of covariance. In a case study of common tern (Sterna hirundo) chicks, in which hatching order is the main determinant of variation in resource acquisition within broods, we find that body mass, but not telomere length or attrition, varies with hatching order. Moreover, body mass and growth positively predict survival to fledging, whereas telomere length and attrition do not. Using a novel statistical method to quantify standardized variance in plasticity, we estimate between‐individual variation in telomere attrition to be only 12% of that of growth. Consistent with the relative invariance of telomere attrition, we find no correlation between age‐specific body mass or growth and telomere attrition. We suggest that common tern chicks prioritize investment in long‐term somatic state (as indicated by canalization of telomere maintenance) over immediate survival benefits of growth as part of an efficient brood reduction strategy that benefits the parents. As such, interspecific variation in the growth–telomere length relationship may be explained by the extent to which parents benefit from rapid mortality of excess offspring.     

Offspring size–number tradeoffs and food quality feedbacks impact population dynamics in a Daphnia–algae system

Population fluctuations can be affected by both extrinsic (e.g. weather patterns, food availability) and intrinsic (e.g. life‐history) factors. A key life‐history tradeoff is the production of offspring size versus number, ranging from many small offspring to few large offspring. Models show that this life‐history tradeoff in offspring size and number, through maturation time, can have significant impacts on population dynamics. However, few manipulative experiments have been conducted that can isolate life‐history effects from impacts of extrinsic factors in consumer–resource systems. We experimentally tested the effect of an offspring size–number tradeoff on population stability and food availability in a consumer–resource system. Using Daphnia pulex, we created a shift from many, small offspring being produced to fewer, larger offspring. Two sets of experiments were performed to examine the interaction of an extrinsic factor (light levels) and intrinsic population structure on dynamics, and we controlled for the ingestion pressure on algal prey at the time of the manipulation. We predicted that the tradeoff would impact internal consumer population characteristics, including biasing the stage structure towards adults, increasing adult size, and increasing average population‐level reproduction. This adult‐dominated stage structure was predicted to then lead to instability and a low quantity–high quality food state. Under all light levels, the manipulated populations became dominated by large adults. Contrary to predictions, the amplitudes of fluctuations in Daphnia biomass were lower in populations shifted to few–large offspring, representing higher stability in these populations. Furthermore, in high light conditions, a stable low Daphnia – high algae biomass (low food quality) state was observed in few–large offspring treatments but not in control (many–small offspring) treatments. Our results show a strong link between light levels as an extrinsic factor and the life‐history tradeoff of consumer offspring size versus number that impacts consumer–resource population dynamics through feedbacks with resource quality.     

Short‐ and long‐term costs of reproduction in a migratory songbird

Costs of reproduction represent a common life‐history trade‐off. Critical to understanding these costs in migratory species is the ability to track individuals across successive stages of the annual cycle. We assessed the effects of total number of offspring fledged and date of breeding completion on pre‐migratory body condition, the schedule of moult and annual survival in a migratory songbird, the Savannah Sparrow Passerculus sandwichensis. Between 2008 and 2010, moult was delayed for individuals that finished breeding later in the breeding period and resulted in reduced lean tissue mass during the pre‐migratory period, suggesting an indirect trade‐off between the timing of breeding completion and condition just prior to migration. Lean tissue mass decreased as the number of offspring fledged increased in 2009, a particularly cool and wet year, illustrating a direct trade‐off between reproductive effort and condition just prior to migration in years when weather is poor. However, using a 17‐year dataset from the same population, we found that parents that fledged young late in the breeding period had the highest survival and that number of offspring fledged did not affect survival, suggesting that individuals do not experience long‐term trade‐offs between reproduction and survival. Taken together, our results suggest that adult Savannah Sparrows pay short‐term costs of reproduction, but that longer‐term costs are mitigated by individual quality, perhaps through individual variation in resource acquisition.     

Experimental evidence of long‐term reproductive costs in a colonial nesting seabird

Trade‐offs between current and future reproduction are central to the evolution of life histories. Experiments that manipulate brood size provide an effective approach to investigating future costs of current reproduction. Most manipulative studies to date, however, have addressed only the short‐term effects of brood size manipulation. Our goal was to determine whether survival or breeding costs of reproduction in a long‐lived species manifest beyond the subsequent breeding season. To this end, we investigated long‐term survival and breeding effects of a multi‐year reproductive cost experiment conducted on black‐legged kittiwakes Rissa tridactyla, a long‐lived colonial nesting seabird. We used multi‐state capture–recapture modeling to assess hypotheses regarding the role of experimentally reduced breeding effort and other factors, including climate phase and colony size and productivity, on future survival and breeding probabilities during the 16‐yr period following the experiment. We found that forced nest failures had a positive effect on breeding probability over time, but had no effect on long‐term survival. This apparent canalization of survival suggests that adult survival is the most important parameter influencing fitness in this long‐lived species, and that adults should pay reproductive costs in ways that do not compromise this critical life history parameter. When declines in adult survival rate are observed, they may indicate populations of conservation concern.     

Effects of variation in resource acquisition during different stages of the life cycle on life‐history traits and trade‐offs in a burying beetle

Individual variation in resource acquisition should have consequences for life‐history traits and trade‐offs between them because such variation determines how many resources can be allocated to different life‐history functions, such as growth, survival and reproduction. Since resource acquisition can vary across an individual's life cycle, the consequences for life‐history traits and trade‐offs may depend on when during the life cycle resources are limited. We tested for differential and/or interactive effects of variation in resource acquisition in the burying beetle Nicrophorus vespilloides. We designed an experiment in which individuals acquired high or low amounts of resources across three stages of the life cycle: larval development, prior to breeding and the onset of breeding in a fully crossed design. Resource acquisition during larval development and prior to breeding affected egg size and offspring survival, respectively. Meanwhile, resource acquisition at the onset of breeding affected size and number of both eggs and offspring. In addition, there were interactive effects between resource acquisition at different stages on egg size and offspring survival. However, only when females acquired few resources at the onset of breeding was there evidence for a trade‐off between offspring size and number. Our results demonstrate that individual variation in resource acquisition during different stages of the life cycle has important consequences for life‐history traits but limited effects on trade‐offs. This suggests that in species that acquire a fixed‐sized resource at the onset of breeding, the size of this resource has larger effects on life‐history trade‐offs than resources acquired at earlier stages.     

Plastic reproductive allocation as a buffer against environmental stochasticity – linking life history and population dynamics to climate

Empirical work suggest that long‐lived organisms have adopted risk sensitive reproductive strategies where individuals trade the amount of resources spent on reproduction versus survival according to expected future environmental conditions. Earlier studies also suggest that climate affects population dynamics both directly by affecting population vital rates and indirectly through long‐term changes in individual life histories. Using a seasonal and state‐dependent individual‐based model we investigated how environmental variability affects the selection of reproductive strategies and their effect on population dynamics. We found that: (1) dynamic, i.e. plastic, reproductive strategies were optimal in a variable climate. (2) Females in poor and unpredictable climatic regimes allocated fewer available resources in reproduction and more in own somatic growth. This resulted in populations with low population densities, and a high average female age and body mass. (3) Strong negative density dependence on offspring body mass and survival, along with co‐variation between climatic severity and population density, resulted in no clear negative climatic effects on reproductive success and offspring body mass. (4) Time series analyses of population growth rates revealed that populations inhabiting benign environments showed the clearest response to climatic perturbations as high population density prohibited an effective buffering of adverse climatic effects as individuals were not able to gain sufficient body reserves during summer. Regularly occurring harsh winters ‘harvested’ populations, resulting in persistent low densities, and released them from negative density dependent effects, resulting in high rewards for a given resource allocation.     

Components of breeding performance in two competing species: habitat heterogeneity, individual quality and density-dependence

Density‐dependent breeding performance due to habitat heterogeneity has been shown to regulate populations of territorial species, since the progressive occupation of low quality territories as breeding density increases may cause a decline in the mean per capita fecundity of a population while variation in fecundity increases. Although the preemptive use of sites may relegate low quality individuals to sites of progressively lower suitability, few studies on density dependence have tried to separate the effects of territory quality from individual quality, and none have simultaneously considered the effects of heterospecific competitors. Using two long‐term monitored populations, we assessed the relative contribution of habitat heterogeneity and bird quality (in terms of age) on the productivity of sympatric golden Aquila chrysaetos and Bonelli's eagles Hieraaetus fasciatus under different scenarios of intra‐ and inter‐specific competition. Productivity (number of offspring fledged) varied among territories and average annual productivity was negatively related to its variability in both species and populations, thus giving some support to the habitat heterogeneity hypothesis. However, the effect of habitat heterogeneity on productivity became non‐significant when parental age and local density estimators were included in multivariate analyses. Therefore, temporal changes in bird quality (age) combined with intra‐ and interspecific competition explained variability in territory productivity rather than habitat heterogeneity among territories per se. The recruitment of subadult breeders, a surrogate of mortality in eagles, strongly varied among territories. Habitat heterogeneity in productivity may thus arise not because sites differ in suitability for reproduction but because of differences in factors affecting survival. Territories associated with high mortality risks have a higher probability of being occupied by young birds, whose lower quality, interacting with the density competitors, leads to a reduction of productivity. Site‐dependent variability in adult survival and interspecific competition may be extensive, but so far largely overlooked, factors to be seriously considered for the site‐dependent population regulation framework.     

The trade‐off between clutch size and egg mass in tree swallows Tachycineta bicolor is modulated by female body mass

Egg production is a costly component of reproduction for female birds in terms of energy expenditure and maternal investment. Because resources are typically limited, clutch size and egg mass are expected to be constrained, and this putative trade‐off between offspring number and size is at the core of life history theory. Nevertheless, empirical evidence for this trade‐off is equivocal at best, as individual heterogeneity in resource acquisition and allocation may hamper the detection of the negative correlation between egg number and mass within populations. Here, we investigated how female body mass and landscape composition influences clutch size, egg mass, and the relationship between these two traits. To do so, we fitted linear mixed models using data from tree swallows Tachycineta bicolor breeding in a network of 400 nestboxes located along a gradient of agricultural intensity between 2004 and 2011. Our dataset comprised 1463 broods for clutch size analyses and 4371 eggs (from 847 broods laid between 2005–2008) for egg mass analyses. Our results showed that agricultural intensity negatively impacted clutch size, but not egg mass nor the relationship between these two traits. Female mass, on the other hand, modulated the trade‐off between clutch size and egg mass. For heavier females, both traits increased jointly, without evidence of a trade‐off. However, for lighter females, there was a clear negative relationship between clutch size and egg mass. This work shows that accounting for individual heterogeneity in body mass allows the detection of a clutch size/egg mass trade‐off that would have remained undetected otherwise. Identifying habitat and individual effects on resource allocation towards reproductive traits may help bridging the gap between predictions from theory and empirical evidence on life history trade‐offs.     

Resource exploitation and relatedness: implications for offspring size variation within broods

Much of the theory on offspring size variation within a brood relies on unequal maternal allocation of resources to each embryo. However, maternal allocation strategies are subject to an inherent conflict between mothers and offspring: individual offspring, being more closely related to themselves than to their siblings, should always prefer a larger share of the available resources than that which is optimal from their mother's perspective. Thus, in species where mothers cannot unilaterally impose a resource allocation strategy, offspring can respond to this conflict by competing for more resources than is maternally optimal. Here we show that variation in offspring size within a brood can arise as a by‐product of competition between siblings over a common resource, even when 1) there are no competitive inequalities within families, and 2) maternal investment per brood is fixed. Moreover, we show that size variance among offspring increases with increasing levels of competition, brought about by decreasing relatedness among siblings. Conflict thus offers a simple, testable and, potentially general, explanation for the wide variability in offspring size seen in nature. This extends explanatory hypotheses for offspring size variation beyond those of maternal effects, under which most explanations have been subsumed to date.     

Effects of developmental conditions on growth,stress and telomeres in black‐legged kittiwake chicks

Early‐life conditions can drive ageing patterns and life history strategies throughout the lifespan. Certain social, genetic and nutritional developmental conditions are more likely to produce high‐quality offspring: those with good likelihood of recruitment and productivity. Here, we call such conditions “favoured states” and explore their relationship with physiological variables during development in a long‐lived seabird, the black‐legged kittiwake (Rissa tridactyla). Two favoured states were experimentally generated by manipulation of food availability and brood size, while hatching order and sex were also explored as naturally generating favoured states. Thus, the favoured states we explored were high food availability, lower levels of sibling competition, hatching first and male sex. We tested the effects of favoured developmental conditions on growth, stress, telomere length (a molecular marker associated with lifespan) and nestling survival. Generation of favoured states through manipulation of both the nutritional and social environments furthered our understanding of their relative contributions to development and phenotype: increased food availability led to larger body size, reduced stress and higher antioxidant status, while lower sibling competition (social environment) led to lower telomere loss and longer telomere lengths in fledglings. Telomere length predicted nestling survival, and wing growth was also positively correlated with telomere length, supporting the idea that telomeres may indicate individual quality, mediated by favoured states.     

Environmental cues influence parental brood structure decisions in the burying beetle <Emphasis Type="Italic">Nicrophorus marginatus</Emphasis>

Parents evaluate multiple extrinsic and intrinsic cues when making decisions associated with reproduction. These decisions often reflect classic trade-offs between the cost of a strategy and its perceived fitness payoff. Life history theory predicts that when parents experience austere conditions, reproductive success is increased by producing fewer but larger offspring with a competitive advantage in this environment. Conversely, parents experiencing favorable conditions are expected to increase current reproductive success by favoring quantity over quality of offspring. We tested the predictions of life history theory using Nicrophorus marginatus (Coleoptera: Silphidae), a burying beetle species that exhibits infanticide during biparental care and hypervariable adult size across populations, by employing a factorial design that manipulated density and nutritional quality of food. We measured (1) the average number of offspring produced, (2) the average individual size of offspring, and (3) the sex ratio of the offspring. We found no effect of density or food quality on offspring sex ratio, but mean offspring size and number differed between low and high-density treatments. Nutritional environment interacted with density effects such that parents with access to high quality diets were able to modulate offspring size and number to match the perceived competitive environment, whereas those in poor nutritional condition appeared to exhibit physiological constraints to producing optimal brood structures.     

Differences in Patterns of Reproductive Allocation between the Sexes in Nicrophorus orbicollis

Organisms are selected to maximize lifetime reproductive success by balancing the costs of current reproduction with costs to future survival and fecundity. Males and females typically face different reproductive costs, which makes comparisons of their reproductive strategies difficult. Burying beetles provide a unique system that allows us to compare the costs of reproduction between the sexes because males and females are capable of raising offspring together or alone and carcass preparation and offspring care represent the majority of reproductive costs for both sexes. Because both sexes perform the same functions of carcass preparation and offspring care, we predict that they would experience similar costs and have similar life history patterns. In this study we assess the cost of reproduction in male Nicrophorus orbicollis and compare to patterns observed in females. We compare the reproductive strategies of single males and females that provided pre- and post-hatching parental care. There is a cost to reproduction for both males and females, but the sexes respond to these costs differently. Females match brood size with carcass size, and thus maximize the lifetime number of offspring on a given size carcass. Males cull proportionately more offspring on all carcass sizes, and thus have a lower lifetime number of offspring compared to females. Females exhibit an adaptive reproductive strategy based on resource availability, but male reproductive strategies are not adaptive in relation to resource availability.